S. Wanless’s research while affiliated with UK Centre for Ecology & Hydrology and other places

Understanding the responses of seabirds to climate-induced variations in phenology and abundance of their prey is key to developing ecosystem-based fisheries management measures that benefit higher trophic levels. The match/mismatch hypothesis (MMH) emphasizes the need to consider synchrony in the seasonal cycles of predators and prey, while the match/mismatch/abundance hypothesis (MMAH) proposes that prey abundance may reinforce/compensate mismatch effects. This study considers the effects of both variations in seasonal availability and abundance of lesser sandeel Ammodytes marinus on hatching, fledging and breeding success of 5 seabird species: black-legged kittiwake Rissa tridactyla , Atlantic puffin Fratercula arctica , razorbill Alca torda , common guillemot Uria aalge and European shag Gulosus aristotelis . Consistent with MMH, temporal asynchrony between sandeel availability and seabird breeding schedules affected productivity in 4 species. The effects of trophic asynchrony were either reinforced or compensated by sandeel abundance for some species, supporting MMAH. Breeding success in the late-breeding kittiwake was high when conditions favoured both high sandeel abundance and temporal synchrony while the cost of asynchrony could be compensated by high sandeel abundance in the earlier-breeding puffin. Differential effects of sandeel abundance and trophic synchrony at different stages of the seabird breeding season suggest that distinct mechanisms are involved. The effects were most evident in the most sandeel-reliant seabirds. As further disruption of sandeel phenology and abundance is anticipated under the current climate crisis, the present study is an important step towards understanding bottom-up effects of environmental change on higher trophic levels.

Supplementary data to "Six pelagic seabird species of the North Atlantic engage in a fly-and-forage strategy during their migratory movements"

Seabirds are undergoing drastic declines globally and spend the non-breeding season at sea, making it challenging to study the drivers of their survival. Harsh weather and changes in climate conditions can have large impacts on seabird population dynamics through increased mortality. The intensity and persistence of extreme events are forecasted to increase with global warming. As shared conditions can induce population synchrony, multi-population studies of key demographic parameters are imperative to explore the influence of climate change. We used long-term mark-recapture data and position data to determine non-breeding stop-over areas of five Atlantic puffin (Fratercula arctica) populations over a latitudinal gradient in the north-eastern Atlantic (56°11’–70°23’N). We investigated synchrony in adult survival in relation to shared stop-over areas. We quantified effects of extreme extra-tropical cyclones (ETCs) specific to populations’ stop-over areas and the North Atlantic Oscillation on adult survival. Populations with overlapping stop-over areas exhibited temporal synchrony in survival rates. Winter ETCs negatively influenced survival in one population, which was the one most exposed to extreme weather, but did not directly influence adult survival in the other four populations. Synchrony among populations with shared stop-over areas highlights the importance of these areas for adult survival, a key life-history rate. However, extreme weather was not identified as a driving factor for four of the populations. This suggests other factors in these areas, likely related to bottom-up trophic interactions, as environmental drivers of synchrony in the survival of Atlantic puffins.

Tracking data of marine predators are increasingly used in marine spatial management. We developed a spatial data set with estimates of the monthly distribution of 6 pelagic seabird species breeding in the Northeast Atlantic. The data set was based on year-round global location sensor (GLS) tracking data of 2356 adult seabirds from 2006-2019 from a network of seabird colonies, data describing the physical environment and data on seabird population sizes. Tracking and environmental data were combined in monthly species distribution models (SDMs). Cross-validations were used to assess the transferability of models between years and breeding locations. The analyses showed that birds from colonies close to each other (<500 km apart) used the same nonbreeding habitats, while birds from distant colonies (>1000 km) used colony-specific and, in many cases, non-overlapping habitats. Based on these results, the SDM from the nearest model colony was used to predict the distribution of all seabird colonies lying within a species-specific cut-off distance (400-500 km). Uncertainties in the predictions were estimated by cluster bootstrap sampling. The resulting data set consisted of 4692 map layers, each layer predicting the densities of birds from a given species, colony and month across the North Atlantic. This data set represents the annual distribution of 23.5 million adult pelagic seabirds, or 87% of the Northeast Atlantic breeding population of the study species. We show how the data set can be used in population and spatial management applications, including the detection of population-specific nonbreeding habitats and identifying populations influenced by marine protected areas.

Bird migration is commonly defined as a seasonal movement between breeding and non-breeding grounds. It generally involves relatively straight and directed large-scale movements, with a latitudinal change, and specific daily activity patterns comprising less or no foraging and more traveling time. Our main objective was to describe how this general definition applies to seabirds. We investigated migration characteristics of 6 pelagic seabird species (little auk Alle alle, Atlantic puffin Fratercula arctica, common guillemot Uria aalge, Brünnich’s guillemot U. lomvia, black-legged kittiwake Rissa tridactyla and northern fulmars Fulmarus glacialis). We analysed an extensive geolocator positional and saltwater immersion dataset from 29 colonies in the North-East Atlantic and across several years (2008−2019). We used a novel method to identify active migration periods based on segmentation of time series of track characteristics (latitude, longitude, net-squared displacement). Additionally, we used the saltwater immersion data of geolocators to infer bird activity. We found that the 6 species had, on average, 3 to 4 migration periods and 2 to 3 distinct stationary areas during the non-breeding season. On average, seabirds spent the winter at lower latitudes than their breeding colonies and followed specific migration routes rather than non-directionally dispersing from their colonies. Differences in daily activity patterns were small between migratory and stationary periods, suggesting that all species continued to forage and rest while migrating, engaging in a ‘fly-and-forage’ migratory strategy. We thereby demonstrate the importance of habitats visited during seabird migrations as those that are not just flown over, but which may be important for re-fuelling.

Species breeding at high latitudes face a significant challenge of surviving the winter. Such conditions are particularly severe for diurnal marine endotherms such as seabirds. A critical question is therefore what behavioural strategies such species adopt to maximise survival probability. We tested 3 hypotheses: (1) they migrate to lower latitudes to exploit longer day length (‘sun-chasing’), (2) they forage at night (‘night-feeding’), or (3) they target high-quality food patches to minimise foraging time (‘feasting’). We studied the winter migration and foraging strategies of European shags Phalacrocorax aristotelis from 6 colonies across a latitudinal gradient from temperate regions to north of the Arctic Circle using geolocators deployed over 11 winters. We found evidence for ‘sun-chasing’, whereby average southerly movements were greatest from colonies at higher latitudes. However, a proportion of individuals from higher latitudes remained resident in winter and, in the absence of daylight, they foraged during twilight and only very occasionally during the night. At lower latitudes, there was little evidence that individuals migrated south, nocturnal feeding was absent, and twilight feeding was infrequent, suggesting that there was sufficient daylight in winter. There was no evidence that winter foraging time was lowest at higher latitudes, as predicted by the ‘feasting’ hypothesis. Our results suggest that shags adopt different behavioural strategies to survive the winter across their latitudinal range, dictated by the differing light constraints. Our study highlights the value of multi-colony studies in testing key hypotheses to explain population persistence in seabird species that occur over large latitudinal ranges.

Sampling seabirds The vastness of the worlds' oceans makes them difficult to monitor. Seabirds that forage and breed across oceans globally have been recognized as sentinels of ocean health. Sydeman et al. looked across seabird species of both the Northern and Southern Hemispheres and found varying patterns. Northern Hemisphere species exhibited greater signs of stress and reduced breeding success, indicative of low fish resources. Southern Hemisphere species showed less impact on reproductive output, suggesting that the fish populations there have thus far been less disturbed. The differences across hemispheres indicate different strategies for conservation, with active recovery needed in the north and enhanced protection in the south. Science , abf1772, this issue p. 980

Migration is a widespread strategy for escaping unfavourable conditions during winter, but the extent to which populations that segregate during the breeding season aggregate during the non-breeding season is poorly understood. Low non-breeding season aggregation may be associated with higher likelihood of overlap with threats, but with fewer populations affected, whereas high aggregation may result in a lower probability of exposure to threats, but higher overall severity. We investigated non-breeding distributions and extent of population aggregation in 2 sympatrically breeding auks. We deployed geolocation-immersion loggers on common guillemots Uria aalge and razorbills Alca torda at 11 colonies around the northern UK and tracked their movements across 2 non-breeding seasons (2017-18 and 2018-19). Using 290 guillemot and 135 razorbill tracks, we mapped population distributions of each species and compared population aggregation during key periods of the non-breeding season (post-breeding moult and mid-winter), observing clear interspecific differences. Razorbills were largely distributed in the North Sea, whereas guillemot distributions were spread throughout Scottish coastal waters and the North, Norwegian and Barents Seas. We found high levels of aggregation in razorbills and a strong tendency for colony-specific distributions in guillemots. Therefore, razorbills are predicted to have a lower likelihood of exposure to marine threats, but more severe potential impact due to the larger number of colonies affected. This interspecific difference may result in divergent population trajectories, despite the species sharing protection at their breeding sites. We highlight the importance of taking whole-year distributions into account in spatial planning to adequately protect migratory species.

Offshore renewable developments (ORDs) may negatively affect seabirds, in particular due to collisions with turbine blades, displacement to less favourable habitats and barrier effects to movement. Many so-called 'sub-lethal effects', whereby individuals birds are not killed instantaneously by an interaction with the wind farm, but behaviour is affected in the short term, may have knock on effects on energetic budgets and, in turn, demographic rates such as survival and productivity. A key potential process linking sub-lethal effects of ORDs and demography is the relationship between adult body condition at the end of the breeding season and survival probability the following winter. However, our understanding of this relationship is limited. Using long-term data collected on the Isle of May, Scotland, this report investigates the relationship between body mass during breeding and survival for 4 seabird species. For puffins, we found evidence for a strongly positive relationship between individual-specific body mass at the end of the breeding season and the probability of survival to the start of the next breeding season. In contrast, in kittiwakes, guillemots and razorbills, the estimated effect sizes were considerably smaller. In conclusion, using the best available UK data and employing advanced methods of statistical analysis, we found evidence for a positive relationship between end-of-breeding season body mass and the survival of puffins with less evidence of an effect in kittiwakes, guillemots and razorbills. We consider these estimates more suitable for use, especially in a UK context, than relationships estimated from other studies.

Identifying drivers of population trends in migratory species is difficult, as they can face many stressors while moving through different areas and environments during the annual cycle. To understand the potential of migrants to adjust to perturbations, it is critical to study the connection of different areas used by different populations during the annual cycle (i.e. migratory connectivity). Using a large-scale tracking data set of 662 individual seabirds from 2 sympatric auk meta-populations (common guillemots Uria aalge and Brünnich’s guillemots U. lomvia ) breeding in 12 colonies throughout the Northeast Atlantic, we estimated migratory connectivity in seasonal space use as well as occupied environmental niches. We found strong migratory connectivity, within and between species. This was apparent through a combination of seasonal space use and occupied environmental niches. Brünnich’s guillemot populations grouped into 2 and common guillemot populations into 5 previously undescribed spatiotemporal clusters. Common guillemot populations clustered in accordance with the variable population trends exhibited by the species, while Brünnich’s guillemot populations are declining everywhere where known within the study area. Individuals from different breeding populations in both species were clustered in their space and environmental use, utilising only a fraction of the potential species-wide range. Further, space use varied among seasons, emphasising the variable constraints faced by both species during the different stages of their annual cycle. Our study highlights that considering spatiotemporal dynamics, not only in space but also in occupied environmental niches, improves our understanding of migratory connectivity and thus population vulnerability in the context of global change.